Effect of different water supply regimes on growth and size hierarchy in spring wheat populations under mulched with clear plastic film
Introduction
Most plant populations consist of relatively few large individuals and many small ones, and the few large individuals account for most of biomass of the population (Obeid et al., 1967, Weiner, 1985, Weiner and Thomas, 1986, Benjamin and Hardwick, 1986). This distribution of biomass has been called “size hierarchies” or “size inequalities” (Weiner and Solbrig, 1984). Due to modem breeding and intensive crop management practices (e.g. genetically uniform pure stands, weed-free, irrigation and fertilizer use, controlled planting date, depth and density), size hierarchies in even-aged plant result mainly from uneven interplant competition within populations (Weiner, 1985, Weiner and Thomas, 1986, Bonan, 1988, Bonan, 1991, Wu and Wang, 1999, Weiner et al., 2001). The ecological and evolutionary significance of size hierarchies is enormous (Harper, 1977, Solbrig, 1980, Wu and Wang, 1999, Xin et al., 1998). For example, the smallest plants suffer density-dependent mortality, and within a population, size appears to be correlated with fitness (Solbrig, 1980, Weiner, 1985).
The arid and semi-arid regions contribute to about 52.5% of the total territory in China. In the region, heat and light is sufficient, Water is principal limit factor in crop production (Wei and Zhao, 1995). There are two main methods to improve water condition of crop growth in the region: mulching and irrigation using rainwater-harvesting.
Crop production in semiarid areas is generally dependent on rainfall, so soil moisture conservation is vital for grain production (Zhao, 1996, Li et al., 1999). Mulching of soil may reduce water loss through evaporation, and therefore may increase water available to plants and grain yield of crops (Cantero-Martinez et al., 1995, O’Leary and Connor, 1997, Li et al., 1999). Studies on vegetable crops have demonstrated that mulches provide several benefits to crop production through soil and water conservation, improved soil physical and chemical properties, and enhanced soil biological activity (Unger, 1975, Tindall et al., 1991). In China's Loess Plateau, plastic film mulching has been utilized in the cultivation of corn (Zea mays L.), cotton (Gossypium hirsutum L.), vegetables and fruit (Zhang and Ma, 1994, Han and Wan, 1995). Recent years this technique has also been introduced in spring wheat. A few studies have been conducted to determine the effect of plastic film mulches on the grain yield potential of spring wheat in the semi-arid regions of Northwest China (Li and Lan, 1995, Li et al., 1999, Du et al., 1999). These studies have shown that mulching increases the number of tillers and grain yield of spring wheat, and mulched spring wheat had a faster seedling emergence and greater dry matter production. Unfortunately, no detailed study has been conducted to describe differences in reproductive allocation between mulched and non-mulched spring wheat populations. This question is critical for determining whether a spring wheat population makes more efficient use of resources under plastic film mulched conditions. In addition, spring wheat plants normally produce tillers, many of which do not survive to produce grain-bearing spikes. Increasing the percentage of tillers that survive has been hypothesized as an approach for increasing yield of wheat plants. But some researchers suggested that as high as 50–70% abortive tillers in certain wheat cultivars (Dewey and Albrechtsen, 1985) is a higher growth redundancy for spring wheat population for higher grain yield (Donald, 1968, Kirby and Jones, 1977).
In recent years, rainwater-harvesting technology has been introduced into dryland agricultural in North China, which improves water availability to the crops (Li et al., 1995, Zhao et al., 1995) and made water control (WC) turning into reality in this areas.
Some important agronomic characters of spring wheat (such as plant height, spike length, grain number per spike, and grain yield) grown along the water control gradient showed that the variation was extensive (Zhu and Shang, 1988). Zi-Zhen et al. (2004) reported that grain yield, spike length, fertile spikelets per spike and grain number per spike increased significantly with irrigation (90 mm) than without irrigation. Among them, grain yield increased 111.7%, spike length increased 54.2%, fertile spikelets per spike increased 71.1% and grain number per spike increased 98.1%. Changing resource pools (e.g. water or nutrient availability) may affect the distribution of biomass (Weiner, 1985, Schmitt et al., 1986, Bonan, 1988, Morse and Bazzaz, 1994, Xin et al., 1998, Wu and Wang, 1999, Duan and Zhao, 1996). Studies of spring wheat populations grown under water limiting conditions showed that the degree of size hierarchies increased with water deficits (Duan and Zhao, 1996, Xin et al., 1998, Wu and Wang, 1999).
The purpose of this study was to investigate (1) whether there was any difference in plant size hierarchy and distributions of spring wheat populations along water control gradient. (2) Whether there was any difference in plant size hierarchy and distribution of spring wheat populations between plastic film mulching and non-mulching. (3) To explore the relationship between size hierarchy and grain yield of spring wheat. (4) To determine the effects of water on the degrees of growth redundancy (increased percentage of abortive tillers and decreased reproductive allocation) in spring wheat population and to explore its mechanism in relation to size hierarchy and life history strategies.
Section snippets
Materials and methods
Field experiments were conducted at the experimental station of Lanzhou University, Gansu province, PR China (35.8°N, 103.7°E, and 1517 m asl). Annual mean precipitation is 316 mm (rain and snow) about half of which occurs between July and September. Monthly mean air temperature and mean rainfall in recent 30 years show that the warm and rainy season occurs in the same period. The soil is a loess-like loam, with a bulk density of 1.5 g cm−3, and a field water holding capacity (FWHC, maximum
Grain yield, yield components, and harvest index (HI)
Along the water control gradient, biological yield (aboveground biomass), grain yield and HI decreased significantly (Table 2). Compared with WC3, biological yields in WC2 and WC1 decreased 6.3, 15.9% in mulching and 9.0, 23.0% in non-mulching, respectively. Grain yields in WC2 and WC1 were lower by 10.0, 31.2% in mulching and 13.4, 32.1% in non-mulching, respectively. The grain yield in mulched treatments was significantly greater (+36.0–41.5%) than that in the non-mulched control (Table 2).
Water control, intraspecific competition and size hierarchy in spring wheat populations
This study suggests that drought stress decreases mean plant biomass, it increases both the relative variation in plant biomass and the concentration of mass within a small fraction of the population. This is supported by earlier studies (Duan and Zhao, 1996, Xin et al., 1998, Wu and Wang, 1999) conducted at fields or in pots.
Increase in size hierarchy may serve as an index of a population's competitive status under environmental stress (Weiner, 1985, Weiner, 1990, Wu and Wang, 1999, Duan and
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